Misplaced Pages

#871128

184-511: Velociraptor ( / v ə ˌ l ɒ s ɪ ˈ r æ p t ər , v ə ˈ l ɒ s ɪ r æ p t ər / ; lit.   ' swift thief ' ) is a genus of small dromaeosaurid dinosaurs that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago . Two species are currently recognized, although others have been assigned in the past. The type species

368-467: A Protoceratops (MPC-D 100/512) and Velociraptor (MPC-D 100/25) fossilized in combat and provides an important window regarding direct evidence of predator-prey behavior in non-avian dinosaurs. In the 1960s and early 1970s, many Polish-Mongolian paleontological expeditions were conducted to the Gobi Desert with the objective of fossil findings. In 1971, the expedition explored several localities of

552-403: A feather covering—have been reported from the ulna of a single Velociraptor specimen (IGM 100/981), which represents an animal of estimated 1.5 m (4.9 ft) long and 15 kg (33 lb) in weight. The spacing of 6 preserved knobs suggests that 8 additional knobs may have been present, giving a total of 14 quill knobs that developed large secondaries ("wing" feathers stemming from

736-557: A species : see Botanical name and Specific name (zoology) . The rules for the scientific names of organisms are laid down in the nomenclature codes , which allow each species a single unique name that, for animals (including protists ), plants (also including algae and fungi ) and prokaryotes ( bacteria and archaea ), is Latin and binomial in form; this contrasts with common or vernacular names , which are non-standardized, can be non-unique, and typically also vary by country and language of usage. Except for viruses ,

920-465: A body mass around 14.1–19.7 kg (31–43 lb). It nevertheless shared many of the same anatomical features. It was a bipedal , feathered carnivore with a long tail and an enlarged sickle-shaped claw on each hindfoot, which is thought to have been used to tackle and restrain prey . Velociraptor can be distinguished from other dromaeosaurids by its long and low skull , with an upturned snout. Velociraptor (commonly referred to as "raptor")

1104-653: A common occurrence in Tugriken Shireh. Since its discovery, the Tugriken Shireh locality has yielded some of the most significant specimens of Protoceratops , such as the Fighting Dinosaurs , in situ individuals—a preservation condition also known as "standing" individuals or specimens in some cases—, authentic nests, and small herd-like groups. Specimens from this locality are usually found in articulation, suggesting possible mass mortality events. Stephan N. F. Spiekman and colleagues reported

1288-409: A decrease in the presence of primitive premaxillary teeth, hence supporting a growing change in the populations. In 1955, paleontologist Georg Haas examined the overall skull shape of Protoceratops and attempted to reconstruct its jaw musculature . He suggested that the large neck frill was likely an attachment site for masticatory muscles. Such placement of the muscles may have helped to anchor

1472-496: A fossilized theropod embryo inside an egg (MPC-D 100/971) from the Djadokhta Formation. They identified this embryo as an oviraptorid dinosaur and the eggshell, upon close examination, turned out be that of elongatoolithid eggs and thereby the oofamily Elongatoolithidae was concluded to represent the eggs of oviraptorids. This find proved that the nest AMNH 6508 belonged to Oviraptor and rather than an egg-thief,

1656-411: A fully or well-developed nasal thermoregulation apparatus as modern endothermic animals do. Norell with colleagues in 1995 reported one V. mongoliensis skull bearing two parallel rows of small punctures on its frontal bones that, upon closer examination, match the spacing and size of Velociraptor teeth. They suggested that the wound was likely inflicted by another Velociraptor during a fight within

1840-427: A key ancestor for the ceratopsid lineage, suggesting that it ultimately led to the evolution of large-bodied ceratopsians such as Styracosaurus and Triceratops . Such lineage was suggested to have started from the primitive ceratopsian Psittacosaurus . He also regarded Protoceratops as one of the first "frilled" ceratopsians to appear in the fossil record. However, in 1975 Maryanska and Osmolska argued that it

2024-403: A large phylogenetic analysis based on skull biomechanical characters—provided by 160 Mesozoic dinosaur species—to analyze the multiple emergences of herbivory among non-avian dinosaurs. Their results found that herbivorous dinosaurs mainly followed two distinct modes of feeding, either processing food in the gut—characterized by relatively gracile skulls and low bite forces —or the mouth, which

SECTION 10

#1732788101872

2208-646: A later date. Maxillae and a lacrimal (the main tooth-bearing bones of the upper jaw, and the bone that forms the anterior margin of the eye socket, respectively) recovered from the Bayan Mandahu Formation in 1999 by the Sino-Belgian Dinosaur Expeditions were found to pertain to Velociraptor , but not to the type species V. mongoliensis . Pascal Godefroit and colleagues named these bones V. osmolskae (for Polish paleontologist Halszka Osmólska ) in 2008. However,

2392-651: A later homonym of a validly published name is a nomen illegitimum or nom. illeg. ; for a full list refer to the International Code of Nomenclature for algae, fungi, and plants and the work cited above by Hawksworth, 2010. In place of the "valid taxon" in zoology, the nearest equivalent in botany is " correct name " or "current name" which can, again, differ or change with alternative taxonomic treatments or new information that results in previously accepted genera being combined or split. Prokaryote and virus codes of nomenclature also exist which serve as

2576-628: A long time and redescribed as new by a range of subsequent workers, or if a range of genera previously considered separate taxa have subsequently been consolidated into one. For example, the World Register of Marine Species presently lists 8 genus-level synonyms for the sperm whale genus Physeter Linnaeus, 1758, and 13 for the bivalve genus Pecten O.F. Müller, 1776. Within the same kingdom, one generic name can apply to one genus only. However, many names have been assigned (usually unintentionally) to two or more different genera. For example,

2760-509: A longer evolutionary history compared to P. andrewsi , or simply a direct descendant of P. andrewsi . The difference in morphologies between Protoceratops also suggests that the nearby Bayan Mandahu Formation is slightly younger than the Djadokhta Formation. In 2020, Czepiński analyzed several long-undescribed protoceratopsid specimens from the Udyn Sayr and Zamyn Khondt localities of the Djadokhta Formation. One specimen (MPC-D 100/551B)

2944-495: A maniraptoran, possibly deinonychosaur taxon. Nevertheless, in 2011 an authentic nest of Protoceratops was reported and described by David E. Fastovsky and colleagues. The nest (MPC-D 100/530) containing 15 articulated juveniles was collected from the Tugriken Shireh locality of the Djadokhta Formation during the work of Mongolian-Japanese paleontological expeditions. Gregory M. Erickson and team in 2017 reported an embryo-bearing egg clutch (MPC-D 100/1021) of Protoceratops from

3128-592: A national treasure of Mongolia, and in 2000 it was loaned to the American Museum of Natural History in New York City for a temporary exhibition. Between 1988 and 1990, a joint Chinese - Canadian team discovered Velociraptor remains in northern China. American scientists returned to Mongolia in 1990, and a joint Mongolian-American expedition to the Gobi, led by the American Museum of Natural History and

3312-497: A paper on their 2008 discovery of shed teeth of what they believed to be a Velociraptor near a tooth-marked jaw bone of what they believed to be a Protoceratops in the Bayan Mandahu Formation. The authors concluded that the find represented "late-stage carcass consumption by Velociraptor " as the predator would have eaten other parts of a freshly killed Protoceratops before biting in the jaw area. The evidence

3496-624: A partial P. andrewsi skull (RGM 818207) in the collections of the Naturalis Biodiversity Center , Netherlands in 2015. Since Protoceratops fossils are only found in the Gobi Desert of Mongolia and this specimen was likely discovered during the Central Asiatic Expeditions, the team concluded that this skull was probably acquired by Delft University between 1940 and 1972 as part of a collection transfer. Protoceratopsid remains were recovered in

3680-814: A possible competition with the more predatory theropods over carcasses , however, as the animal tissue ingestion was occasional and not the bulk of their diet, the energy flow in ecosystems was relatively simple. You Hailu and Peter Dodson in 2004 suggested that the premaxillary teeth of Protoceratops may have been useful for selective cropping and feeding. In 2009, Kyo Tanque and team suggested that basal ceratopsians, such as protoceratopsids, were most likely low browsers due to their relatively small body size. This low-browsing method would have allowed to feed on foliage and fruits within range, and large basal ceratopsians may have consumed tougher seeds or plant material not available to smaller basal ceratopsians. David J. Button and Lindsay E. Zanno in 2019 performed

3864-520: A rare occurrence in dinosaurs. During maturation, the skull and neck frill underwent rapid growth. Protoceratops were hunted by Velociraptor , and one particularly famous specimen (the Fighting Dinosaurs ) preserves a pair of them locked in combat. Protoceratops used to be characterized as nocturnal because of the large sclerotic ring around the eye, but they are now thought to have been cathemeral (active at dawn and dusk). In 1900 Henry Fairfield Osborn suggested that Central Asia may have been

SECTION 20

#1732788101872

4048-409: A reference for designating currently accepted genus names as opposed to others which may be either reduced to synonymy, or, in the case of prokaryotes, relegated to a status of "names without standing in prokaryotic nomenclature". An available (zoological) or validly published (botanical) name that has been historically applied to a genus but is not regarded as the accepted (current/valid) name for

4232-405: A rigid structure. The neck was rather short and had poor flexibility. The atlas was the smallest cervical and consisted mainly of the centrum because the neural arch (upper, and pointy vertebral region) was a thin, narrow bar of bone that extended upwards and backward to the base of the axis neural spine . The capitular facet (attachment site for chevrons ; also known as cervical ribs)

4416-592: A sandstorm or a collapsed dune . During the Third Central Asiatic Expedition in 1923, a nearly complete Protoceratops skeleton (specimen AMNH 6418) was collected at the Flaming Cliffs. Unlike other specimens, it was discovered in a rolled-up position with its skull preserving a thin, hard, and wrinkled layer of matrix (surrounding sediments ). This specimen was later described in 1940 by Brown and Schlaikjer, who discussed

4600-433: A similar range of grasping motion. The short metatarsus and foot strength, however, would have been more similar to that of owls . The RPR method of predation would be consistent with other aspects of Velociraptor ' s anatomy, such as their unusual jaw and arm morphology. The arms, which could exert a lot of force but were likely covered in long feathers, may have been used as flapping stabilizers for balance while atop

4784-407: A single row that created a shearing surface. Both dentary and maxillary teeth presented marked homodonty —a dental condition where the teeth share a similar shape and size. P. andrewsi bore two small, peg to spike-like teeth that were located on the underside of each premaxilla. The second premaxillary tooth was larger than the first one. Unlike dentary and maxillary teeth, the premaxillary dentition

4968-410: A small hole near its posterior end, called surangular foramen or fenestra. Both bones were the largest elements of the lower jaw of Velociraptor , contributing to virtually its entire length. Below them were the smaller splenial and angular , closely articulated to each other. The articular , located on the inner side of the surangular, was a small element that joined the quadrate of

5152-484: A sort of 'wastebin' taxon, where many unrelated species were grouped together). As dinosaur discoveries multiplied, Velociraptor was later recognized as a dromaeosaurid. All dromaeosaurids have also been referred to the family Archaeopterygidae by at least one author (which would, in effect, make Velociraptor a flightless bird). In the past, other dromaeosaurid species, including Deinonychus antirrhopus and Saurornitholestes langstoni , have sometimes been classified in

5336-407: A stocky and strongly T-shaped bone. As a whole, the orbit or orbital fenestra (eye socket)—formed by the lacrimal, jugal, frontal, and postorbital—was large and near circular in shape, being longer than taller. When seen from above, a pair of large and markedly rounded holes were present near the rear of the skull (the temporal fenestrae), whose main components were the postorbital and squamosal. Behind

5520-443: A struggle. The specimen, nicknamed the "Fighting Dinosaurs", has been examined and studied by numerous researchers and paleontologists, and there are various opinions on how the animals were buried and preserved altogether. Though a drowning scenario has been proposed by Barsbold, such a hypothesis is considered unlikely given the arid paleoenvironments of the Djadokhta Formation. It is generally thought that they were buried alive by

5704-471: A struggling prey animal, along with the stiff counterbalancing tail. The jaws, thought by Fowler and colleagues to be comparatively weak, would have been useful for row saw motion bites like the modern day Komodo dragon , which also has a weak bite, to finish off its prey if the kicks were not powerful enough. These predatory adaptations working together may also have implications for the origin of flapping in paravians . In 2010, Hone and colleagues published

Velociraptor - Misplaced Pages Continue

5888-427: A taxon; however, the names published in suppressed works are made unavailable via the relevant Opinion dealing with the work in question. In botany, similar concepts exist but with different labels. The botanical equivalent of zoology's "available name" is a validly published name . An invalidly published name is a nomen invalidum or nom. inval. ; a rejected name is a nomen rejiciendum or nom. rej. ;

6072-455: A total of c. 520,000 published names (including synonyms) as at end 2019, increasing at some 2,500 published generic names per year. "Official" registers of taxon names at all ranks, including genera, exist for a few groups only such as viruses and prokaryotes, while for others there are compendia with no "official" standing such as Index Fungorum for fungi, Index Nominum Algarum and AlgaeBase for algae, Index Nominum Genericorum and

6256-477: Is V. mongoliensis , named and described in 1924. Fossils of this species have been discovered in the Djadochta Formation , Mongolia . A second species, V. osmolskae , was named in 2008 for skull material from the Bayan Mandahu Formation , China . Smaller than other dromaeosaurids like Deinonychus and Achillobator , Velociraptor was about 1.5–2.07 m (4.9–6.8 ft) long with

6440-429: Is derived from Greek hellenikos (meaning Greek) and rhis (meaning nose) in reference to its broad and angular snout, which is reminiscent of the straight profiles of Greek sculptures . In 2017 abundant protoceratopsid material was reported from Alxa near Bayan Mandahu, and it may be preferable to P. hellenikorhinus . Viktor Tereshchenko and Vladimir R. Alifanov in 2003 named a new protoceratopsid dinosaur from

6624-612: Is discouraged by both the International Code of Zoological Nomenclature and the International Code of Nomenclature for algae, fungi, and plants , there are some five thousand such names in use in more than one kingdom. For instance, A list of generic homonyms (with their authorities), including both available (validly published) and selected unavailable names, has been compiled by the Interim Register of Marine and Nonmarine Genera (IRMNG). The type genus forms

6808-462: Is evidence that the ancestors of dromaeosaurids could fly, making Velociraptor and other large members of this family secondarily flightless, though it is possible the large wing feathers inferred in the ancestors of Velociraptor had a purpose other than flight. The feathers of the flightless Velociraptor may have been used for display, for covering their nests while brooding, or for added speed and thrust when running up inclined slopes. Because of

6992-427: Is in honor of Andrews for his prominent leadership during the expeditions. They identified Protoceratops as an ornithischian dinosaur closely related to ceratopsians representing a possible common ancestor between ankylosaurs and ceratopsians . Since Protoceratops was more primitive than any other known ceratopsian at that time, Granger and Gregory coined the new family Protoceratopsidae , mostly characterized by

7176-615: Is in tribute to the Polish paleontologist Roman Kozłowski . They also named the new genus and species of protoceratopsid Bagaceratops rozhdestvenskyi , known from specimens of the nearby Hermiin Tsav locality. In 1990 the Russian paleontologist Sergei Mikhailovich Kurzanov referred additional material from Hermiin Tsav to P. kozlowskii . However, he noted that there were enough differences between P. andrewsi and P. kozlowskii , and erected

7360-612: Is now thought that the animals were buried in sand, either from a collapsing dune or in a sandstorm . Burial must have been extremely rapid, judging from the lifelike poses in which the animals were preserved. Parts of the Protoceratops are missing, which has been seen as evidence of scavenging by other animals. Comparisons between the scleral rings of Velociraptor , Protoceratops , and modern birds and reptiles indicates that Velociraptor may have been nocturnal , while Protoceratops may have been cathemeral , active throughout

7544-486: Is one of the dinosaur genera most familiar to the general public due to its prominent role in the Jurassic Park films. In reality, however, Velociraptor was roughly the size of a turkey , considerably smaller than the approximately 2 m (6.6 ft) tall and 90 kg (200 lb) reptiles seen in the novels and films (which were based on members of the related genus Deinonychus ). Today, Velociraptor

Velociraptor - Misplaced Pages Continue

7728-533: Is only limited fossil evidence to support this theory for dromaeosaurids in general and none specific to Velociraptor itself. Dromeosaur footprints in China suggest that a few other raptor genera may have hunted in packs, but there have been no conclusive examples of pack behavior found. In 2011, Denver Fowler and colleagues suggested a new method by which dromaeosaurs like Velociraptor and similar dromaeosaurs may have captured and restrained prey. This model, known as

7912-418: Is similar to dromaeosaurids in anatomy, feather type, bone structure and even the narrow anatomy of the nasal passages (usually a key indicator of metabolism). The kiwi is a highly active, if specialized, flightless bird, with a stable body temperature and a fairly low resting metabolic rate, making it a good model for the metabolism of primitive birds and dromaeosaurids. In 2023, Seishiro Tada and team examined

8096-460: Is somewhat arbitrary. Although all species within a genus are supposed to be "similar", there are no objective criteria for grouping species into genera. There is much debate among zoologists about whether enormous, species-rich genera should be maintained, as it is extremely difficult to come up with identification keys or even character sets that distinguish all species. Hence, many taxonomists argue in favor of breaking down large genera. For instance,

8280-474: Is the type species , and the generic name is permanently associated with the type specimen of its type species. Should the specimen turn out to be assignable to another genus, the generic name linked to it becomes a junior synonym and the remaining taxa in the former genus need to be reassessed. In zoological usage, taxonomic names, including those of genera, are classified as "available" or "unavailable". Available names are those published in accordance with

8464-431: Is the obtained cladogram , showing the position of Protoceratops and Bagaceratops : Graciliceratops Bagaceratops Protoceratops Zuniceratops Turanoceratops Ceratopsidae Leptoceratopsidae Longrich and team in 2010 indicated that highly derived morphology of P. hellenikorhinus —when compared to P. andrewsi —indicates that this species may represent a lineage of Protoceratops that had

8648-411: Is very unlikely that protoceratopsids evolved from psittacosaurids , and also unlikely that they gave rise to the highly derived (advanced) ceratopsids. The first point was supported by the numerous anatomical differences between protoceratopsids and psittacosaurids, most notably the extreme reduction of some hand digits in the latter group—a trait much less pronounced in protoceratopsids. The second point

8832-547: Is well known to paleontologists , with over a dozen described fossil skeletons. One particularly famous specimen preserves a Velociraptor locked in combat with a Protoceratops . During an American Museum of Natural History expedition to the Flaming Cliffs (Bayn Dzak or Bayanzag) of the Djadochta Formation , Gobi Desert , on 11 August 1923, Peter Kaisen discovered the first Velociraptor fossil known to science—a crushed but complete skull, associated with one of

9016-427: The frontals (bones of the skull roof ). Both parietals were coossified (fused), creating a long ridge on the center of the frill. The jugal was deep and sharply developed and along with the quadratojugal they formed a horn-like extension that pointed to below at the lateral sides of the skull. The epijugal (tip region of the jugal) was separated from the jugal by a prominent suture ; this suture

9200-477: The Barun Goyot Formation , noting numerous similarities with Protoceratops . Even though their respective skull anatomy had substantial differences, their postcranial skeleton was virtually the same. The phylogenetic analysis performed by the team recovered both protoceratopsids as sister taxa, indicating that Bagaceratops and Protoceratops were anatomically and systematically related. Below

9384-467: The Cold War , expeditions by Soviet and Polish scientists, in collaboration with Mongolian colleagues, recovered several more specimens of Velociraptor . The most famous is part of the " Fighting Dinosaurs " specimen ( MPC-D 100/25; formerly IGM, GIN, or GI SPS), discovered by a Polish-Mongolian team in 1971. The fossil preserves a Velociraptor in battle against a Protoceratops . It is considered

SECTION 50

#1732788101872

9568-621: The International Code of Zoological Nomenclature ; the earliest such name for any taxon (for example, a genus) should then be selected as the " valid " (i.e., current or accepted) name for the taxon in question. Consequently, there will be more available names than valid names at any point in time; which names are currently in use depending on the judgement of taxonomists in either combining taxa described under multiple names, or splitting taxa which may bring available names previously treated as synonyms back into use. "Unavailable" names in zoology comprise names that either were not published according to

9752-824: The International Plant Names Index for plants in general, and ferns through angiosperms, respectively, and Nomenclator Zoologicus and the Index to Organism Names for zoological names. Totals for both "all names" and estimates for "accepted names" as held in the Interim Register of Marine and Nonmarine Genera (IRMNG) are broken down further in the publication by Rees et al., 2020 cited above. The accepted names estimates are as follows, broken down by kingdom: The cited ranges of uncertainty arise because IRMNG lists "uncertain" names (not researched therein) in addition to known "accepted" names;

9936-535: The Mongolian Academy of Sciences , turned up several well-preserved skeletons. One such specimen, MPC-D 100/980, was nicknamed "Ichabodcraniosaurus" by Norell's team because the fairly complete specimen was found without its skull (an allusion to the Washington Irving character Ichabod Crane ). While Norell and Makovicky provisionally considered it a specimen of Velociraptor mongoliensis , it

10120-490: The holotype specimen (AMNH 6251) of Protoceratops —in reddish sandstones . It was subsequently analyzed by the paleontologist Walter W. Granger who identified it as reptilian . On 21 September, the expedition returned to Beijing, and even though it was set up to look for remains of human ancestors, the team collected numerous dinosaur fossils and thus provided insights into the rich fossil record of Asia. Back in Beijing,

10304-447: The leaves . This feeding method was likely more efficient in protoceratopsids as the enamel surface of Protoceratops was coarsely-textured and the tips of the micro-serrations developed on the basis of the teeth, probably helping to crumble vegetation. Based on their respective peg-like shape and reduced microornamentation, Dauphin and colleagues suggested that the premaxillary teeth of Protoceratops had no specific function. In 1991,

10488-400: The occipital condyle : a rounded and bulbous protuberance that meet the first vertebra of the neck. The lower jaw of Velociraptor comprised mainly the dentary, splenial, angular, surangular, and articular bones. The dentary was a very long, weakly curved, and narrow element that developed several alveoli on its top surface. On its posterior end, it meet the surangular . It had

10672-637: The phylogenetic analysis conducted by James G. Napoli and team in 2021 during the description of Kuru , showing the position of Velociraptor : Saurornitholestes [REDACTED] Bambiraptor [REDACTED] Achillobator [REDACTED] Utahraptor [REDACTED] Dromaeosaurus [REDACTED] Linheraptor [REDACTED] Tsaagan [REDACTED] Deinonychus [REDACTED] Adasaurus [REDACTED] Kuru [REDACTED] Balaur [REDACTED] Shri [REDACTED] Velociraptor [REDACTED] In 2007 Alan H. Turner and colleagues reported

10856-404: The platypus belongs to the genus Ornithorhynchus although George Shaw named it Platypus in 1799 (these two names are thus synonyms ) . However, the name Platypus had already been given to a group of ambrosia beetles by Johann Friedrich Wilhelm Herbst in 1793. A name that means two different things is a homonym . Since beetles and platypuses are both members of the kingdom Animalia,

11040-405: The "raptor prey restraint" (RPR) model of predation, proposes that dromaeosaurs killed their prey in a manner very similar to extant accipitrid birds of prey : by leaping onto their quarry, pinning it under their body weight, and gripping it tightly with the large, sickle-shaped claws. These researchers proposed that, like accipitrids, the dromaeosaur would then begin to feed on the animal while it

11224-491: The 1970s from the Khulsan locality of the Barun Goyot Formation , Mongolia, during the work of several Polish-Mongolian paleontological expeditions. In 1975, Polish paleontologists Teresa Maryańska and Halszka Osmólska described a second species of Protoceratops which they named P. kozlowskii . This new species was based on the Khulsan material, mostly consisting of juvenile skull specimens. The specific name, kozlowskii ,

SECTION 60

#1732788101872

11408-505: The 2010s, including expanded versions of the analyses that found support for Velociraptorinae, have failed to resolve it as a distinct group, but rather have suggested it is a paraphyletic grade which gave rise to the Dromaeosaurinae . When first described in 1924, Velociraptor was placed in the family Megalosauridae , as was the case with most carnivorous dinosaurs at the time (Megalosauridae, like Megalosaurus , functioned as

11592-490: The 2013 study noted that while "the elongate shape of the maxilla in V. osmolskae is similar to that of V. mongoliensis ," phylogenetic analysis found it to be closer to Linheraptor , making the genus paraphyletic ; thus, V. osmolskae might not actually belong to the genus Velociraptor and requires reassessment. Paleontologists Mark A. Norell and Peter J. Makovicky in 1997 described new and well preserved specimens of V. mongoliensis , namely MPC-D 100/985 collected from

11776-483: The Bayan Mandahu locality of the Bayan Mandahu Formation , Inner Mongolia, in 1995 and 1996 during Sino -Belgian paleontological expeditions. The holotype (IMM 95BM1/1) and paratype (IMM 96BM1/4) specimens consist of large skulls lacking body remains. The holotype skull was found facing upwards, a pose that has been reported in Protoceratops specimens from Tugriken Shireh. The specific name, hellenikorhinus ,

11960-551: The Bayn Dzak locality, Bainoceratops efremovi . This genus was based on a few dorsal (back) vertebrae that were stated to differ from those of Protoceratops . In 2006 North American paleontologists Peter Makovicky and Mark A. Norell suggested that Bainoceratops may be synonymous with Protoceratops as most of the traits used to separate the former from the latter have been reported from other ceratopsians including Protoceratops itself, and they are more likely to fall within

12144-558: The Djadokhta and Nemegt formations. During fieldwork on 3 August several fossils of Protoceratops and Velociraptor were found at the Tugriken Shire locality (Djadokhta Formation) including a block containing one of each. The individuals in this block were identified as a P. andrewsi and V. mongoliensis . Although the conditions surrounding their burial were not fully understood, it was clear that they died simultaneously in

12328-431: The Flaming Cliffs yet again. During this year more eggs and nests were collected, alongside well-preserved and complete specimens of Protoceratops . By this time, Protoceratops had become one of the most abundant dinosaurs of the region with more than 100 specimens known, including skulls and skeletons of multiple individuals at different growth stages. Though more remains of Protoceratops were collected in later years of

12512-473: The French botanist Joseph Pitton de Tournefort (1656–1708) is considered "the founder of the modern concept of genera". The scientific name (or the scientific epithet) of a genus is also called the generic name ; in modern style guides and science, it is always capitalised. It plays a fundamental role in binomial nomenclature , the system of naming organisms , where it is combined with the scientific name of

12696-754: The Russian paleontologist Konstantin E. Mikhailov named the new oogenus Protoceratopsidovum from the Barun Goyot and Djadokhta formations, with the type species P. sincerum and additional P. fluxuosum and P. minimum . This ootaxon was firmly stated as belonging to protoceratopsid dinosaurs since they were the predominant dinosaurs where the eggs were found and some skeletons of Protoceratops were found in close proximity to Protoceratopsidovum eggs. More specifically, Mikhailov stated that P. sincerum and P. minimum were laid by Protoceratops , and P. fluxuosum by Breviceratops . However, also during 1994, Norell and colleagues reported and briefly described

12880-433: The Third Central Asiatic Expedition of 1923, Andrews and team discovered the holotype specimen of Oviraptor in association with some of the first known fossilized dinosaur eggs (nest AMNH 6508), in the Djadokhta Formation. Each egg was elongated and hard-shelled, and due to the proximity and high abundance of Protoceratops in the formation , these eggs were believed at the time to belong to this dinosaur. This resulted in

13064-606: The Tugrik Shireh locality in 1993, and MPC-D 100/986 collected in 1993 from the Chimney Buttes locality. The team briefly mentioned another specimen, MPC-D 100/982, which by the time of this publication remained undescribed. In 1999 Norell and Makovicky provided more insights into the anatomy of Velociraptor with additional specimens. Among these, MPC-D 100/982 was partially described and figured, and referred to V. mongoliensis mainly based on cranial similarities with

13248-460: The also fossiliferous Ukhaa Tolgod locality, discovered during paleontological expeditions of the American Museum of Natural History and Mongolian Academy of Sciences . This clutch comprises at least 12 eggs and embryos with only 6 embryos preserving nearly complete skeletons. Norell with colleagues in 2020 examined fossilized remains around the eggs of this clutch which indicate a soft-shelled composition. The Fighting Dinosaurs specimen preserves

13432-844: The anatomy of P. andrewsi in extensive detail using newly prepared specimens from the Asiatic expeditions. In 1963, the Mongolian paleontologist Demberelyin Dashzeveg reported the discovery of a new fossiliferous locality of the Djadokhta Formation: Tugriken Shireh. Like the neighbouring Bayn Dzak, this new locality contained an abundance of Protoceratops fossils. During the 1960s to 1970s, Polish-Mongolian and Russian-Mongolian paleontological expeditions collected new, partial to complete specimens of Protoceratops at this locality, making this dinosaur species

13616-420: The anterior one) and their size became smaller towards the end. The caudal vertebrae decreased in size progressively towards the end and had very elongated neural spines in the mid-series, forming a sail -like structure. This elongation started from the first to the fourteenth caudal. The centra were heterocoelous (saddle-shaped at both facets). On the anterior caudals they were broad, however, from

13800-458: The axial skeleton of this specimen. Protoceratops was a relatively small-sized ceratopsian , with both P. andrewsi and P. hellenikorhinus estimated up to 2–2.5 m (6.6–8.2 ft) in length, and around 62–104 kg (137–229 lb) in body mass. Although similar in overall body size, the latter had a relatively greater skull length. Both species can be differentiated by the following characteristics: The skull of Protoceratops

13984-442: The base for higher taxonomic ranks, such as the family name Canidae ("Canids") based on Canis . However, this does not typically ascend more than one or two levels: the order to which dogs and wolves belong is Carnivora ("Carnivores"). The numbers of either accepted, or all published genus names is not known precisely; Rees et al., 2020 estimate that approximately 310,000 accepted names (valid taxa) may exist, out of

14168-416: The bony core of the claw is preserved. The thick abdominal wall of skin and muscle of large prey species would have been difficult to slash without a specialized cutting surface. The slashing hypothesis was tested during a 2005 BBC documentary, The Truth About Killer Dinosaurs . The producers of the program created an artificial Velociraptor leg with a sickle claw and used a pork belly to simulate

14352-525: The capacity to walk around bipedally if necessary . They were characterized by a proportionally large skull , short and stiff neck, and neck frill . The frill was likely used for display or intraspecific combat , as well as protection of the neck and anchoring of jaw muscles. A horn-like structure was present over the nose, which varied from a single structure in P. andrewsi to a double, paired structure in P. hellenikorhinus . The "horn" and frill were highly variable in shape and size across individuals of

14536-534: The center of origin of most animal species, including humans , which caught the attention of explorer and zoologist Roy Chapman Andrews . This idea later gave rise to the First (1916 to 1917), Second (1919) and Third (1921 to 1930) Central Asiatic Expeditions to China and Mongolia , organized by the American Museum of Natural History under the direction of Osborn and field leadership of Andrews. The team of

14720-481: The coexistence and sympatric (altogether) evolution of both Bagaceratops and Protoceratops at this one locality; (2) the rise of B. rozhdestvenskyi in a different region and eventual migration to Udyn Sayr; (3) hybridization between the two protoceratopsids given the near placement of both Bayan Mandahu and Djadokhta; (4) anagenetic (progressive evolution) evolutionary transition from P. andrewsi to B. rozhdestvenskyi . Among scenarios, an anagenetic transition

14904-427: The coronoid process of the dentary, being obscured by the jugal. The surangular was near triangular in shape and in old individuals it was coossified together with the coronoid process. The angular was located below the two latter bones and behind the dentary. It was a large and somewhat rounded bone that complemented the curvature of the dentary. On its inner surface it was attached to the articular . The articular

15088-474: The day during short intervals, suggesting that the fight may have occurred at twilight or during low-light conditions. The distinctive claw, on the second digit of dromaeosaurids, has traditionally been depicted as a slashing weapon; its assumed use being to cut and disembowel prey. In the "Fighting Dinosaurs" specimen, the Velociraptor lies underneath, with one of its sickle claws apparently embedded in

15272-495: The diagnostic (identifier) features used to distinguish these taxa to be largely present in Bagaceratops and thus becoming synonyms of this genus. Under this reasoning, Protoceratopsidae consists of Bagaceratops , Breviceratops , and Protoceratops . Below are the proposed relationships among Protoceratopsidae by Czepiński: In 2019 Bitnara Kim and colleagues described a relatively well-preserved Bagaceratops skeleton from

15456-461: The dinosaur's prey. Though the sickle claw did penetrate the abdominal wall, it was unable to tear it open, indicating that the claw was not used to disembowel prey. Remains of Deinonychus , a closely related dromaeosaurid, have commonly been found in aggregations of several individuals. Deinonychus has also been found in association with the large ornithopod Tenontosaurus , which has been cited as evidence of cooperative (pack) hunting. However,

15640-470: The expeditions, they were most abundant in the 1922 to 1925 seasons. Gregory and Charles C. Mook published another description of Protoceratops in 1925, discussing its anatomy and relationships. Thanks to the large collection of skulls found in the expeditions, they concluded that Protoceratops represented a ceratopsian more primitive than ceratopsids and not an ankylosaur-ceratopsian ancestor. In 1940, Barnum Brown and Erich Maren Schlaikjer described

15824-423: The forearm). However, the specimen number has been corrected to IGM 100/3503 and its referral to Velociraptor may require reevaluation, pending further study. Nevertheless, there is strong phylogenetic evidence from other dromaeosaurid relatives that indicates the presence of feathers in Velociraptor , including dromaeosaurids such as Daurlong , Microraptor , or Zhenyuanlong . The skull of Velociraptor

16008-446: The form "author, year" in zoology, and "standard abbreviated author name" in botany. Thus in the examples above, the genus Canis would be cited in full as " Canis Linnaeus, 1758" (zoological usage), while Hibiscus , also first established by Linnaeus but in 1753, is simply " Hibiscus L." (botanical usage). Each genus should have a designated type , although in practice there is a backlog of older names without one. In zoology, this

16192-534: The fourth to the ninth all cervicals were relatively equal in size and proportions. Their neural spines were smaller than the first three vertebrae and the development of the capitular facet diminished from the fourth cervical onwards. The dorsal vertebrae were similar in shape and size. Their neural spines were elongated and sub-rectangular in shape with a tendency to become more elongated in posterior vertebrae. The centra were large and predominantly amphiplatian (flat on both facets) and circular when seen from

16376-414: The frill varied by individual; some had short, compact frills, while others had frills nearly half the length of the skull. The squamosal touched the jugal (cheekbone) and was very enlarged and high having a curved end that built the borders of the frill. The parietals were the posteriormost bones of the skull and major elements of the frill. In a top view they had a triangular shape and were joined by

16560-403: The front. Sometimes in old individuals the last dorsal vertebra was somewhat coosified to the first sacral. The sacral vertebrae were firmly coosified giving form to the sacrum, which was connected to the inner sides of both ilia. Their neural spines were broad, not coosified, and rather consistent in length. The centra were mainly opisthocoelous (concave on the posterior facet and convex on

16744-425: The front. The premaxilla had 4 alveoli (meaning that 4 teeth were developed), and the maxilla had 11 alveoli. At the dentary, between 14–15 alveoli were present. All teeth present at the premaxilla were poorly curved, and the two first teeth were the longest, with the second having a characteristic large size. The maxillary teeth were more slender, recurved, and most notably, the lower end was strongly more serrated than

16928-737: The generic name (or its abbreviated form) still forms the leading portion of the scientific name, for example, Canis lupus lupus for the Eurasian wolf subspecies, or as a botanical example, Hibiscus arnottianus ssp. immaculatus . Also, as visible in the above examples, the Latinised portions of the scientific names of genera and their included species (and infraspecies, where applicable) are, by convention, written in italics . The scientific names of virus species are descriptive, not binomial in form, and may or may not incorporate an indication of their containing genus; for example,

17112-492: The genus Velociraptor . Since Velociraptor was the first to be named, these species were renamed Velociraptor antirrhopus and V. langstoni . As of 2008, the only currently recognized species of Velociraptor are V. mongoliensis and V. osmolskae . However, several studies have found " V. " osmolskae to be distantly related to V. mongoliensis . Below are the results for the Eudromaeosauria phylogeny based on

17296-519: The holotype skull, although they stated that differences were present between the pelvic region of this specimen and other Velociraptor specimens. This relatively well-preserved specimen including the skull was discovered and collected in 1995 at the Bayn Dzak locality (specifically at the "Volcano" sub-locality). Martin Kundrát in a 2004 abstract compared the neurocranium of MPC-D 100/982 to another Velociraptor specimen, MPC-D 100/976. He concluded that

17480-418: The holotype was actually a mature individual that perished brooding the eggs. Moreover, phylogenetic analyses published in 2008 by Darla K. Zelenitsky and François Therrien have shown that Protoceratopsidovum represents the eggs of a maniraptoran more derived than oviraptorids and not Protoceratops . The description of the eggshell of Protoceratopsidovum has further confirmed that they in fact belong to

17664-525: The humerus, radius, and ulna. The humerus (upper arm bone) was large and slender, and at the lower part it met with both radius and ulna. The radius had a slightly recurved shape and was longer than the ulna. A concavity was present on its upper part, serving as the connection with the humerus and forming the elbow. The ulna was a rather short bone with a straight shape. The manus (hand) of Protoceratops had five digits (fingers). The first three fingers had unguals (claw bones) and were

17848-432: The idea that a newly defined genus should fulfill these three criteria to be descriptively useful: Moreover, genera should be composed of phylogenetic units of the same kind as other (analogous) genera. The term "genus" comes from Latin genus , a noun form cognate with gignere ('to bear; to give birth to'). The Swedish taxonomist Carl Linnaeus popularized its use in his 1753 Species Plantarum , but

18032-439: The importance of the Protoceratops finds, and the genus was hailed as the "long-sought ancestor of Triceratops ". Most fossils were in an excellent state of preservation with even sclerotic rings (delicate ocular bones) preserved in some specimens, quickly making Protoceratops one of the best-known dinosaurs from Asia. After spending much of 1924 making plans for the next fieldwork seasons, in 1925 Andrews and team explored

18216-401: The individual. The forward facing and closely located orbits combined with a narrow snout, gave Protoceratops a well-developed binocular vision . Behind the eye was a slightly smaller fenestra known as the infratemporal fenestra , formed by the curves of the jugal and squamosal. The last openings of the skull were two parietal fenestrae (holes in the frill). The lower jaw of Protoceratops

18400-559: The interpretation of the contemporary Oviraptor as an egg predatory animal, an interpretation also reflected in its generic name. In 1975 , the Chinese paleontologist Zhao Zikui named the new oogenera Elongatoolithus and Macroolithus , including them in a new oofamily : the Elongatoolithidae . As the name implies, they represent elongated dinosaur eggs, including some of referred ones to Protoceratops . In 1994

18584-550: The interpretations proposing direct relationships with more derived ceratopsians unsupported. In 2019 Czepiński analyzed a vast majority of referred specimens to the ceratopsians Bagaceratops and Breviceratops , and concluded that most were in fact specimens of the former. Although the genera Gobiceratops , Lamaceratops , Magnirostris , and Platyceratops , were long considered valid and distinct taxa, and sometimes placed within Protoceratopsidae, Czepiński found

18768-417: The jugal, an inverted T-shaped bone (also seen in other dromaeosaurids), known as the quadratojugal , was developed. While the upper end of the quadratojugal joined the squamosal , an irregularly-shaped element, its inner side meet the quadrate . The latter was of great importance for the articulation with the lower jaw. The posteriormost bone was the occipital bone and its projection

18952-412: The lack of horns. The co-authors also agreed with Osborn that Asia, if more thoroughly explored, could solve many major evolutionary gaps in the fossil record. Although not stated in the original description, the generic name , Protoceratops , is intended to mean "first horned face" as it was believed that Protoceratops represented an early ancestor of ceratopsids . Other researchers immediately noted

19136-423: The large dromaeosurine Achillobator is a unique exception to Asian taxa with its deep maxilla. Manabu Sakamoto in 2022 performed a Bayesian phylogenetic predictive modelling framework for estimating jaw muscle parameters and bite forces of several extinct archosaurs, based on skull widths and phylogenetic relationships between groups. Among studied taxa, Velociraptor was scored with a bite force of 304 N , which

19320-853: The larger P. hellenikorhinus . The former was described in 1923 with fossils from the Mongolian Djadokhta Formation , and the latter in 2001 with fossils from the Chinese Bayan Mandahu Formation . Protoceratops was initially believed to be an ancestor of ankylosaurians and larger ceratopsians, such as Triceratops and relatives, until the discoveries of other protoceratopsids. Populations of P. andrewsi may have evolved into Bagaceratops through anagenesis . Protoceratops were small ceratopsians, up to 2–2.5 m (6.6–8.2 ft) long and around 62–104 kg (137–229 lb) in body mass. While adults were largely quadrupedal , juveniles had

19504-558: The larger family Dromaeosauridae. It is often placed within its own subfamily, Velociraptorinae . In phylogenetic taxonomy , Velociraptorinae is usually defined as "all dromaeosaurs more closely related to Velociraptor than to Dromaeosaurus ." However, dromaeosaurid classification is highly variable. Originally, the subfamily Velociraptorinae was erected solely to contain Velociraptor . Other analyses have often included other genera, usually Deinonychus and Saurornitholestes , and more recently Tsaagan . Several studies published during

19688-633: The largest component, with 23,236 ± 5,379 accepted genus names, of which 20,845 ± 4,494 are angiosperms (superclass Angiospermae). By comparison, the 2018 annual edition of the Catalogue of Life (estimated >90% complete, for extant species in the main) contains currently 175,363 "accepted" genus names for 1,744,204 living and 59,284 extinct species, also including genus names only (no species) for some groups. The number of species in genera varies considerably among taxonomic groups. For instance, among (non-avian) reptiles , which have about 1180 genera,

19872-437: The largest digits. The last two were devoid of unguals and had a small size, mostly vestigial (retained, but without important function). Both hand and feet unguals were flat, blunt and hoof-like. The pelvic girdle was formed by the ilium , pubis , and ischium . The ilium was a large element, having a narrow preacetabular process (anterior end) and a wide postacetabular process (posterior end). The pubis

20056-432: The lateral sides, except for the series of caudals. The first five cervical ribs (sometimes called chevrons) were some of the shortest ribs, and among them the first two were longer than the rest. The third to the sixth dorsal (thoracic) ribs were the longest ribs in the skeleton of Protoceratops , the following ribs became smaller in size as they progressed toward the end of the vertebral column. The two last dorsal ribs were

20240-419: The lateral surface of the dentary that connected the coronoid process —a bony projection that extends upwards from the upper surface of the lower jaw behind the tooth row—and surangular. It bore up to 12–14 alveoli on its top margin. Both predentary and dentary had a series of foramina (small pits), the latter mostly on its anterior end. The coronoid (highest point of the lower jaw) was blunt-shaped and touched by

20424-567: The lizard genus Anolis has been suggested to be broken down into 8 or so different genera which would bring its ~400 species to smaller, more manageable subsets. Protoceratops Protoceratops ( / ˌ p r oʊ t oʊ ˈ s ɛr ə t ɒ p s / ; lit.   ' first horned face ' ) is a genus of small protoceratopsid dinosaurs that lived in Asia during the Late Cretaceous , around 75 to 71 million years ago. The genus Protoceratops includes two species: P. andrewsi and

20608-465: The lower jaws, useful for feeding. Yannicke Dauphin and colleagues in 1988 described the enamel microstructure of Protoceratops , observing a non-prismatic outer layer. They concluded that enamel shape does not relate to the diet or function of the teeth as most animals do not necessarily use teeth to process food. The maxillary teeth of ceratopsians were usually packed into a dental battery that formed vertical shearing blades which probably chopped

20792-431: The maxilla formed (predominantly) the antorbital fenestra, one of the several large holes in the skull. Both premaxilla and maxilla had several alveoli ( tooth sockets) on their bottom surfaces. Above the maxilla and making contact with the premaxilla, there was the nasal bone. It was a thin/narrow and elongated bone contributing to the top surface of the snout. Together, both premaxilla and nasal bones gave form to

20976-591: The maxilla of MPC-D 100/982 to strongly differ from specimens referred to Velociraptor . They indicated that this specimen, based on these results, represents a different species. In 2021 Powers with team used Principal Component Analysis to separate dromaeosaurid maxillae, most notably finding that MPC-D 100/982 falls outside the instraspecific variability of V. mongoliensis , arguing for a distinct species. They considered that both V. mongoliensis and this new species were ecologically separated based on their skull anatomy. The team in another 2021 abstract reinforced again

21160-439: The maxilla to be a reliable reference when inferring the shape of the premaxilla and overall snout . For instance, most Asian species have elongated snouts based on the maxilla (namely velociraptorines ), indicating a selective feeding in Velociraptor and relatives, such as picking up small, fast prey. In contrast, most North American eudromaeosaurs, mostly dromaeosaurines, feature a robust and deep maxillar morphology. However,

21344-403: The most (>300) have only 1 species, ~360 have between 2 and 4 species, 260 have 5–10 species, ~200 have 11–50 species, and only 27 genera have more than 50 species. However, some insect genera such as the bee genera Lasioglossum and Andrena have over 1000 species each. The largest flowering plant genus, Astragalus , contains over 3,000 species. Which species are assigned to a genus

21528-428: The name could not be used for both. Johann Friedrich Blumenbach published the replacement name Ornithorhynchus in 1800. However, a genus in one kingdom is allowed to bear a scientific name that is in use as a generic name (or the name of a taxon in another rank) in a kingdom that is governed by a different nomenclature code. Names with the same form but applying to different taxa are called "homonyms". Although this

21712-437: The naris or narial fenestra (nostril opening), which was relatively large and circular. The posterior end of the nasal was joined by the frontal and lacrimal bones. The back or anterior region of the skull was built by the frontal, lacrimal, postorbital, jugal, parietal, quadrate, and quadratojugal bones. The frontal was large element, having a vaguely rectangular shape when seen from above. On its posterior end, this bone

21896-556: The nasal cavities of ectotherm (cold-blooded) or endotherm (warm-blooded) species, in order to evaluate the thermoregulatory physiology of non-avian dinosaurs compared to these groups. They found that the size of the nasal cavity relative to the head size of extant endotherms is larger than those of extant ectotherms, and among taxa, Velociraptor was recovered below the extant endotherms level by reconstructing its nasal respiratory cavity. Tada with team suggested that Velociraptor and most other non-avian dinosaurs may not have possessed

22080-400: The nasal, the main body of the premaxilla touched the maxilla. The maxilla was nearly triangular in shape and the largest element of the snout. On its center or main body, there was a depression developing a small oval to circular-shaped hole, called maxillary fenestra. Though in front of this fenestra were two small openings, referred to as promaxillary fenestrae. The posterior border of

22264-486: The nature of the matrix portion. They stated that this layer had a very skin-like texture and covered mostly the left side of the skull from the snout to the neck frill . Brown and Schlaikjer discarded the idea of possible skin impressions as this skin-like layer was likely a product of the decay and burial of the individual, making the sediments become highly attached to the skull. The potential importance of these remains were unrecognized or given attention, and by 2020

22448-470: The new genus and combination Breviceratops kozlowskii . Though Breviceratops has been regarded as a synonym and juvenile stage of Bagaceratops , Łukasz Czepiński in 2019 concluded that the former has enough anatomical differences to be considered as a separate taxon . In 2001 Oliver Lambert with colleagues named a new and distinct species of Protoceratops , P. hellenikorhinus . The first known remains of P. hellenikorhinus were collected from

22632-523: The only solid evidence for social behavior of any kind among dromaeosaurids comes from a Chinese trackway which shows six individuals of a large species moving as a group. Although many isolated fossils of Velociraptor have been found in Mongolia, none were closely associated with other individuals. Therefore, while Velociraptor is commonly depicted as a pack hunter , as in Jurassic Park , there

22816-505: The orbits, frontals, and lacrimals suffered a shrinkage in relative size as the animal aged; the top border of the nostrils became more vertical; the nasal bones progressively became elongated and narrowed; and the neck frill as a whole also increases in size with age. The neck frill specifically, underwent a dramatic change from a small, flat, and almost rounded structure in juveniles to a large, fan-like one in fully mature Protoceratops individuals. In 2001, Lambert and colleagues considered

23000-545: The overall morphology of the former was more derived (advanced) than the latter, suggesting that they could represent distinct taxa. Mark J. Powers in his 2020 master thesis fully described MPC-D 100/982, which he concluded to represent a new and third species of Velociraptor . This species, which he considered distinct, was stated to mainly differ from other Velociraptor species in having a shallow maxilla morphology. Powers and colleagues also in 2020 used morphometric analyses to compare several dromaeosaurid maxillae, and found

23184-459: The paleontologist Gregory S. Paul stated that contrary to the popular view of ornithischians as obligate herbivores , some groups may have been opportunistic meat-eaters , including the members of Ceratopsidae and Protoceratopsidae. He pointed out that their prominent parrot-like beaks and shearing teeth along with powerful muscles on the jaws suggest an omnivore diet instead, much like pigs, hogs , boars and entelodonts . Such scenario indicates

23368-436: The palpebral protruded upwards from the prefrontal , just above the orbit and slightly meeting the frontal, creating a small horn-like structure. The lacrimal was a near-rectangular bone located in front of the orbit, contributing to the shape of the latter. The sclerotic ring (structure that supports the eyeball ), found inside the orbit, was circular in shape and formed by consecutive bony plates. The snout

23552-495: The presence of another dromaeosaurid in Ukhaa Tolgod, Tsaagan , Napoli and team have noted that the referral of this specimen to Velociraptor is currently subject to reexamination. Examinations of the endocranium of Velociraptor indicate that it was able to detect and hear a wide range of sound frequencies (2,368–3,965 Hz) and could track prey with ease as a result. The endocranium examinations also further cemented

23736-593: The presence of six quill knobs in the ulna of a referred Velociraptor specimen (IGM 100/981) from the Ukhaa Tolgod locality of the Djadochta Formation . Turner and colleagues interpreted the presence of feathers on Velociraptor as evidence against the idea that the larger, flightless maniraptorans lost their feathers secondarily due to larger body size. Furthermore, they noted that quill knobs are almost never found in flightless bird species today, and that their presence in Velociraptor (presumed to have been flightless due to its relatively large size and short forelimbs)

23920-541: The provisions of the ICZN Code, e.g., incorrect original or subsequent spellings, names published only in a thesis, and generic names published after 1930 with no type species indicated. According to "Glossary" section of the zoological Code, suppressed names (per published "Opinions" of the International Commission of Zoological Nomenclature) remain available but cannot be used as the valid name for

24104-463: The raptorial second toe claws ( AMNH 6515). In 1924, museum president Henry Fairfield Osborn designated the skull and claw (which he assumed to come from the hand) as the type specimen of his new genus, Velociraptor . This name is derived from the Latin words velox ('swift') and raptor ('robber' or 'plunderer') and refers to the animal's cursorial nature and carnivorous diet. Osborn named

24288-420: The re-examinations of Turanoceratops in 2009 and Zuniceratops —two critical ceratopsian taxa regarding the evolutionary history of ceratopsids—in 2010 it was concluded that the origin of ceratopsids is unrelated to, and older than the fossil record of Protoceratops and relatives. In most recent/modern phylogenetic analyses Protoceratops and Bagaceratops are commonly recovered as sister taxa , leaving

24472-434: The same species, but there is no evidence of sexual dimorphism . They had a prominent parrot-like beak at the tip of the jaws. P. andrewsi had a pair of cylindrical, blunt teeth near the tip of the upper jaw. The forelimbs had five fingers of which only the first three bore wide and flat unguals . The feet were wide and had four toes with flattened, shovel-like unguals, which would have been useful for digging through

24656-474: The same year have revealed that the right side of the skull remains intact, retaining much of this layer and pending further analysis. Also from the context of the Polish-Mongolian paleontological expeditions, in 1965 an articulated subadult Protoceratops skeleton (specimen ZPAL Mg D-II/3) was collected from the Bayn Dzak locality of the Djadokhta Formation. In the 2000s during the preparation of

24840-503: The sand. The hindlimbs were longer than the forelimbs. The tail was long and had an enigmatic sail -like structure, which may have been used for display, swimming , or metabolic reasons. Protoceratops , like many other ceratopsians, were herbivores equipped with prominent jaws and teeth suited for chopping foliage and other plant material. They are thought to have lived in highly sociable groups of mixed ages. They appear to have cared for their young . They laid soft-shelled eggs ,

25024-407: The scapulae were wide. At their lower region, the scapulae meet the coracoids. The coracoids were relatively elliptical, and sometimes coosified (fused) to the scapulae. The clavicle of Protoceratops was an U to slightly V-shaped element that joined to the upper border of the scapulocoracoid. In its general form, the forelimbs of Protoceratops were shorted than the hindlimbs, and composed by

25208-442: The skull Shackelford had found was sent to the American Museum of Natural History for further study, after which Osborn reached out to Andrews and team via cable, notifying them about the importance of the specimen. In 1923 the expedition again prospected the Flaming Cliffs, this time discovering even more specimens of Protoceratops and also the first remains of Oviraptor , Saurornithoides and Velociraptor . Most notably,

25392-451: The skull morphology and bite forces of various dromaeosaurids, it was discovered that Velociraptor had high bite force resistance compared to other dromaeosaurids such as Dromaeosaurus itself and Deinonychus , the latter of which was much larger. It is theorized by the authors that high bite force resistance was an adaptation towards obtaining food through scavenging more often than through active predation in Velociraptor . Velociraptor

25576-428: The smallest, and the last of them was in contact with the internal surfaces of the ilium. Most of the sacral ribs were fused into the sacrum, and had a rather curved shape. The pectoral girdle of Protoceratops was formed by the scapulocoracoid (fusion of the coracoid and scapula) and clavicle. The scapulae (shoulder blades) were relatively large and rounded on their inner sides. At their upper region,

25760-432: The smallest, while the other elements were of similar shape and length. Protoceratops was in 1923 placed within the newly named family Protoceratopsidae as the representative species by Granger and Gregory. This family was characterized by their overall primitive morphology in comparison to the more derived Ceratopsidae , such as lack of well-developed horn cores and relative smaller body size. Protoceratops itself

25944-699: The species . Because its bone structure shows no sign of healing near the bite wounds and the overall specimen was not scavenged, this individual was likely killed by this fatal wound. In 2001 Molnar and team noted that this specimen is MPC-D 100/976 hailing from the Tugrik Shireh locality, which has also yielded the Fighting Dinosaurs specimen. Genus The composition of a genus is determined by taxonomists . The standards for genus classification are not strictly codified, so different authorities often produce different classifications for genera. There are some general practices used, however, including

26128-425: The species-level separation, noting that additional differences can be found in the hindlimbs. Velociraptor was a small to medium-sized dromaeosaurid , with adults measuring between 1.5–2.07 m (4.9–6.8 ft) long, approximately 0.5 m (1.6 ft) high at the hips, and weighing about 14.1–19.7 kg (31–43 lb). Prominent quill knobs —attachment site of " wing " feathers and direct indicator of

26312-497: The specific name particular to the wolf. A botanical example would be Hibiscus arnottianus , a particular species of the genus Hibiscus native to Hawaii. The specific name is written in lower-case and may be followed by subspecies names in zoology or a variety of infraspecific names in botany . When the generic name is already known from context, it may be shortened to its initial letter, for example, C. lupus in place of Canis lupus . Where species are further subdivided,

26496-478: The specimen has already been completely prepared losing all traces of this skin-like layer. Some elements were damaged in the process such as the rostrum . In 2022 Phil R. Bell and colleagues briefly described these potential soft tissues based on the photographs provided by Brown and Schlaikjer, as well as other ceratopsian soft tissues. However, although the initial perception was that the entire skin-like layer had been removed, photographs shared by Czepiński during

26680-568: The specimen, a fossilized cast of a four-toed digitigrade footprint was found below the pelvic girdle. This footprint was described in 2012 by Grzegorz Niedźwiedzki and colleagues who considered it to represent one of the first reported finds of a dinosaur footprint in association with an articulated skeleton, and also the first one reported for Protoceratops . The limb elements of the skeleton of ZPAL Mg D-II/3 were described in 2019 by paleontologists Justyna Słowiak, Victor S. Tereshchenko and Łucja Fostowicz-Frelik. Tereshchenko in 2021 fully described

26864-412: The standard format for a species name comprises the generic name, indicating the genus to which the species belongs, followed by the specific epithet, which (within that genus) is unique to the species. For example, the gray wolf 's scientific name is Canis lupus , with Canis ( Latin for 'dog') being the generic name shared by the wolf's close relatives and lupus (Latin for 'wolf') being

27048-404: The tail. These were once thought to fully stiffen the tail, forcing the entire tail to act as a single rod-like unit. However, at least one specimen has preserved a series of intact tail vertebrae curved sideways into an S -shape, suggesting that there was considerably more horizontal flexibility than once thought. Velociraptor is a member of the group Eudromaeosauria , a derived sub-group of

27232-403: The taxon is termed a synonym ; some authors also include unavailable names in lists of synonyms as well as available names, such as misspellings, names previously published without fulfilling all of the requirements of the relevant nomenclatural code, and rejected or suppressed names. A particular genus name may have zero to many synonyms, the latter case generally if the genus has been known for

27416-437: The team discovered the first fossilized dinosaur eggs near the holotype of Oviraptor and given how abundant Protoceratops was, the nest was attributed to this taxon . This would later result in the interpretation of Oviraptor as an egg-thief . In the same year, Granger and William K. Gregory formally described the new genus and species Protoceratops andrewsi based on the holotype skull. The specific name , andrewsi ,

27600-521: The theory that the dromaeosaur was an agile, swift predator. Fossil evidence suggesting Velociraptor scavenged also indicates that it was an opportunistic and actively predatory animal, feeding on carrion during times of drought or famine, if in poor health, or depending on the animal's age. In 2020, Powers and colleagues re-examined the maxillae of several eudromaeosaur taxa concluding that most Asian and North American eudromaeosaurs were separated by snout morphology and ecological strategies. They found

27784-537: The third expedition arrived in Beijing in 1921 for the final preparations and started working in the field in 1922. During late 1922 the expedition explored the famous Flaming Cliffs of the Shabarakh Usu region of the Djadokhta Formation , Gobi Desert , now known as the Bayn Dzak region. On 2 September, the photographer James B. Shackelford discovered a partial juvenile skull—which would become

27968-460: The throat of its prey, while the beak of Protoceratops is clamped down upon the right forelimb of its attacker. This suggests Velociraptor may have used its sickle claw to pierce vital organs of the throat, such as the jugular vein , carotid artery , or trachea (windpipe), rather than slashing the abdomen. The inside edge of the claw was rounded and not unusually sharp, which may have precluded any sort of cutting or slashing action, although only

28152-425: The twenty-fifth onwards the centra became elongated alongside the neural spines. On the underside of the caudal vertebrae a series of chevrons were attached, giving form to the lower part of the tail. The first chevron was located at the union of the third and fourth caudals. Chevrons three to nine were the largest and from the tenth onwards they became smaller. All vertebrae of Protoceratops had ribs attached on

28336-400: The type species V. mongoliensis after its country of origin. Earlier that year, Osborn had informally mentioned the animal in a popular press article, under the name "Ovoraptor djadochtari" (not to be confused with the similarly named Oviraptor ), eventually changed into V. mongoliensis during its formal description. While North American teams were shut out of communist Mongolia during

28520-406: The upper one. The arm of Velociraptor was formed by the humerus (upper arm bone), radius and ulna (forearm bones), and manus (hand). Velociraptor , like other dromaeosaurids, had a large manus with three elongated digits (fingers), which ended up in strongly curved unguals (claw bones) that were similar in construction and flexibility to the wing bones of modern birds . The second digit

28704-412: The upper skull, enabling the articulation with the lower jaw. An elongated, near oval-shaped hole was developed in the center of the lower jaw (the mandibular fenestra), and it was produced by the joint of the dentary, surangular, and angular bones. The teeth of Velociraptor were fairly homodont (equal in shape) and had several denticles (serrations), each more strongly serrated on the back edge than

28888-576: The values quoted are the mean of "accepted" names alone (all "uncertain" names treated as unaccepted) and "accepted + uncertain" names (all "uncertain" names treated as accepted), with the associated range of uncertainty indicating these two extremes. Within Animalia, the largest phylum is Arthropoda , with 151,697 ± 33,160 accepted genus names, of which 114,387 ± 27,654 are insects (class Insecta). Within Plantae, Tracheophyta (vascular plants) make up

29072-429: The virus species " Salmonid herpesvirus 1 ", " Salmonid herpesvirus 2 " and " Salmonid herpesvirus 3 " are all within the genus Salmonivirus ; however, the genus to which the species with the formal names " Everglades virus " and " Ross River virus " are assigned is Alphavirus . As with scientific names at other ranks, in all groups other than viruses, names of genera may be cited with their authorities, typically in

29256-418: The wide intraspecific variation range of the concurring P. andrewsi . The authors Brenda J. Chinnery and Jhon R. Horner in 2007 during their description of Cerasinops stated that Bainoceratops , along with other dubious genera, was determined to be either a variant or immature specimen of other genera. Based on this reasoning, they excluded Bainoceratops from their phylogenetic analysis. As part of

29440-410: Was warm-blooded to some degree, as it required a significant amount of energy to hunt. Modern animals that possess feathery or furry coats, like Velociraptor did, tend to be warm-blooded, since these coverings function as insulation. However, bone growth rates in dromaeosaurids and some early birds suggest a more moderate metabolism , compared with most modern warm-blooded mammals and birds. The kiwi

29624-404: Was a large element composed of the predentary , dentary , coronoid , angular and surangular . The predentary (frontmost bone) was very pointed and elongated, having a V-shaped symphyseal (bone union) region at the front. The dentary (teeth-bearing bone) was robust, deep, slightly recurved, and fused to the angular and surangular. A large and thick ridge ran along

29808-436: Was a small dewclaw . The second digit, for which Velociraptor is most famous, was highly modified and held retracted off the ground, which caused Velociraptor and other dromaeosaurids to walk on only their third and fourth digits. It bore a relatively large, sickle-shaped claw, typical of dromaeosaurid and troodontid dinosaurs. This enlarged claw, which could grow to over 6.5 cm (2.6 in) long around its outer edge,

29992-400: Was a smaller bone and had a concavity on its inner surface for the articulation with the quadrate. Protoceratops had leaf-shaped dentary and maxillary teeth that bore several denticles (serrations) on their respective edges. The crowns (upper exposed part) had two faces or lobes that were divided by a central ridge-like structure (also called "primary ridge"). The teeth were packed into

30176-418: Was best supported by Czepiński given the fact that no definitive B. rozhdestvenskyi fossils are found in Udyn Sayr, as expected from a hybridization event; MPC-D 100/551B lacks a well-developed accessory antorbital fenestra (hole behind the nostril openings), a trait expected to be present if B. rozhdestvenskyi had migrated to the area; and many specimens of P. andrewsi recovered at Udyn Sayr already feature

30360-540: Was characterized by features associated with extensive processing such as high bite forces and robust jaw musculature. Ceratopsians (including protoceratopsids), along with Euoplocephalus , Hungarosaurus , parkosaurid , ornithopod and heterodontosaurine dinosaurs, were found to be in the former category, indicating that Protoceratops and relatives had strong bite forces and relied mostly on its jaws to process food. Brown and Schlaikjer in 1940 upon their large description and revision of Protoceratops remarked that

30544-466: Was considered by the authors to be somehow related to ankylosaurians based on skull traits, with a more intensified degree to Triceratops and relatives. Gregory and Charles C. Mook in 1925 upon a more deeper analysis of Protoceratops and its overall morphology, concluded that this taxon represents a ceratopsian more primitive than ceratopsids and not an ankylosaur-ceratopsian ancestor. In 1951 Edwin H. Colbert considered Protoceratops to represent

30728-447: Was devoid of denticles, having a relatively smooth surface. All teeth had a single root (lower part inserted in the alveoli). The vertebral column of Protoceratops had nine cervical (neck), 12 dorsal (back), eight sacral (pelvic) and over 40 caudal (tail) vertebrae. The centra (centrum; body of the vertebrae) of the first three cervicals were coossified together ( atlas , axis and third cervical respectively) creating

30912-535: Was explained on the basis of the already derived anatomy in protoceratopsids like Bagaceratops or Protoceratops (such as the jaw morphology). Maryanska and Osmolska also emphasized that some early members of the Ceratopsidae reflect a much older evolutionary history. In 1998, paleontologist Paul Sereno formally defined Protoceratopsidae as the branch -based clade including all coronosaurs closer to Protoceratops than to Triceratops . Furthermore, with

31096-420: Was formed by a low projection located near the base of the neural arch. The anterior facet of the atlas centrum was highly concave for the articulation of the occipital condyle of the skull. The neural arch and spine of the axis were notably larger than the atlas itself and any other cervical. The axial neural spine was broad and backward developed being slightly connected to that of the third cervical. From

31280-476: Was formed by the nasal , maxilla r, premaxilla r and rostral bones. The nasal was generally rounded but some individuals had a sharp nasal boss (a feature that has been called "nasal horn"). In P. hellenikorhinus this boss was divided in two sharp and long ridges. The maxilla was very deep and had up to 15 alveoli ( tooth sockets) on its underside or teeth bearing surface. The premaxilla had two alveoli on its lower edge—a character that

31464-406: Was in contact with the parietal , and such elements were the main bodies of the skull roof . The lacrimal was a T-shaped bone and its main body was thin and delicated. Its lower end meet the jugal (often called cheek bone), which was a large, sub-triangular-shaped element. Its lower border was notably straight/horizontal. The postorbital was located just above the jugal:

31648-441: Was lower than that of other dromaeosaurids such as Dromaeosaurus (885 N) or Deinonychus (706 N). The " Fighting Dinosaurs " specimen, found in 1971, preserves a Velociraptor mongoliensis and Protoceratops andrewsi in combat and provides direct evidence of predatory behavior. When originally reported, it was hypothesized that the two animals drowned. However, as the animals were preserved in ancient sand dune deposits, it

31832-411: Was more noticeable in adults. The surfaces around the epijugal were coarse, indicating that it was covered by a horny sheath. Unlike the much derived ceratopsids , the frontal and postorbital bones of Protoceratops were flat and lacked horn cores or supraorbital horns. The palpebral (small spur-like bone) joined the prefrontal over the front of the orbit (eye socket). In P. hellenikorhinus

32016-400: Was most likely a predatory device used to restrain struggling prey. As in other dromaeosaurs, Velociraptor tails had prezygapophyses (long bony projections) on the upper surfaces of the vertebrae , as well as ossified tendons underneath. The prezygapophyses began on the tenth tail (caudal) vertebra and extended forward to brace four to ten additional vertebrae, depending on position in

32200-671: Was named as a new species Shri devi in 2021. In 1999, Rinchen Barsbold and Halszka Osmólska reported a juvenile Velociraptor specimen (GIN or IGM 100/2000), represented by a complete skeleton including the skull of a young individual. It was found at the Tugriken Shireh locality of the Djadochta Formation during the context of the Mongolian-Japanese Palaeontological Expeditions. The coauthors stated that detailed descriptions of this and other specimens would be published at

32384-538: Was present at least on P. andrewsi . The rostral bone was devoid of teeth, high and triangular in shape. It had a sharp end and rough texture, which reflects that a rhamphotheca (horny beak ) was present. As a whole, the skull had four pairs of fenestrae (skull openings). The foremost hole, the nares (nostril opening), was oval-shaped and considerably smaller than the nostrils seen in ceratopsids. Protoceratops had large orbits, which measured around 5 cm (50 mm) in diameter and had irregular shapes depending on

32568-447: Was rather elongated and grew up to 23 cm (9.1 in) long. It was uniquely up-curved at the snout region, concave on the upper surface, and convex on the lower surface. The snout, which occupied about 60% of the entire skull length, was notably narrow and mainly formed by the nasal, premaxilla, and maxilla bones. The premaxilla was the anteriormost bone in the skull, and it was longer than taller. While its posterior end joined

32752-461: Was relatively large compared to its body and robustly built. The skull of the type species, P. andrewsi , had an average total length of nearly 50 cm (500 mm). On the other hand P. hellenikorhinus had a total skull length of about 70 cm (700 mm). The rear of the skull gave form to a pronounced neck frill (also known as "parietal frill") mostly composed of the parietal and squamosal bones. The exact size and shape of

32936-432: Was robust and had a rather rounded and pronounced greater trochanter , which was slightly recurved into the inner sides. The tibia (shinbone) was long and slender with a wide lower end. On its upper region a concavity was developed for the joint with the smaller fibula . The pes (foot) were composed of four metatarsal and four toes which bore shovel-like pedal unguals. The first metatarsal and toe were

33120-404: Was seen as supporting the inference from the "Fighting Dinosaurs" fossil that Protoceratops was part of the diet of Velociraptor . In 2012, Hone and colleagues published a paper that described a Velociraptor specimen with a long bone of an azhdarchid pterosaur in its gut. This was interpreted as showing scavenging behaviour. In a 2024 study by Tse, Miller, and Pittman et al., focusing on

33304-490: Was shown to present skull traits that are intermediate between Bagaceratops rozhdestvenskyi (which is native to adjacent Bayan Mandahu and Barun Goyot ) and P. andrewsi . The specimen hails from the Udyn Sayr locality, where Protoceratops remains are dominant, and given the lack of more conclusive anatomical traits, Czepiński assigned the specimen as Bagaceratops sp. He explained that the presence of this Bagaceratops specimen in such unusual locality could be solved by: (1)

33488-451: Was still alive, and prey death would eventually result from blood loss and organ failure. This proposal is based primarily on comparisons between the morphology and proportions of the feet and legs of dromaeosaurs to several groups of extant birds of prey with known predatory behaviors. Fowler found that the feet and legs of dromaeosaurs most closely resemble those of eagles and hawks , especially in terms of having an enlarged second claw and

33672-509: Was the longest of the three digits present, while the first was shortest. The structure of the carpal (wrist) bones prevented pronation of the wrist and forced the manus to be held with the palmar surface facing inward ( medially ), not downward. The pes (foot) anatomy of Velociraptor consisted of the metatarsus—a large element composed of three metatarsals of which the first one was extremely reduced in size—and four digits that developed large unguals. The first digit, as in other theropods,

33856-419: Was the smallest element of the pelvic girdle and it had an irregular shape, although its lower end was developed into a pointed bony projection downward. The ischium was the longest bone of the pelvic girdle. It had an elongated shaft with a somewhat wide lower end. The hindlimbs of Protoceratops were rather long, with a slighter longer tibia (lower leg bone) than femur (thigh bone). The femur (thighbone)

#871128